"This book with contributions from scientists and experts studying stingless bees and meliponiculture gives an insight into pot-pollen which supports the stingless bees, and its many interesting biological characteristics that are only now beginning to be understood by scientists." (Bees for Development Journal, Issue 130, March, 2019)

Forewords

Introduction

Acknowledgements

SECTION 1

Pollen and the Evolution of Mutualism

1. Pot-Pollen as a Discipline. What Does it Include?

1.1. Pot-Pollen and Palynology from an Ecological Point of View

1.2. A Modern Synthesis of Bee-Pollen and Pot-Pollen Study

1.3. Plant Reproduction

1.4. Pollination

1.5. Pollen Biology and Palynology

1.6. Applied Pollen Taxonomy

2. Are Stingless Bees a Broadly Polylectic Group? An Empirical Study of the Adjustments Required for an Improved Assessment of Pollen Diet in Bees

2.1. Introduction

2.2. Pollen Specialization Categories in Bees

2.3. Pollen Analysis of Samples

2.4. Adjustment Calculations to Assess Pollen Specialization Categories in Stingless Bees

2.4.1. Modifying the Number of Foraged Resource Items: Threshold Values and Pollen Type Versus Pollen Species

2.4.2. Modifying the Number of Available Resources: Spatial and Temporal Adjustments

2.5. The Importance of an Appropriate Assessment of Pollen Specialization in Bees: Factors Causing Low Number of Foraged Items

2.5.1 Abundant Versus Minor Pollen Types

2.5.2 Recruitment Behaviour

2.5.3 Intra-nest Pollen Analysis

2.6. Factors Causing High Number of Available Items<

2.7. Polylecty, Broad Polylecty or Simply degrees of polylecty?<

3. Pollen collected by stingless bees: a contribution to understand Amazonian biodiversity

3.1. Introduction

3.1.1 Origin and Evolution of Plant-Bee Interactions

3.2. The Use of Pollen Analysis in the Study of Bees in the Amazon Rainforest

3.3. Diversity of Plants, Stingless Bees and their Interactions in Central Amazon

3.4. Amazonian Bee Diet, Biology and Suggested Interactions Potentially Leading to Pollination

3.5. How to Improve Meliponiculture for Sustainable Development in the Amazon

3.6. Conclusions

4. The Stingless Honey Bees (Apidae, Apinae: Meliponini) in Panama, and Ecology from Pollen Analysis

4.1. An Introduction to the Stingless Honey Bees and Pot-Pollen, in Panama

4.2. Pollen niche, relative specialization, and pollen spectrum

4.2.1 Qualitative and quantitative analyses

4.2.2 Field bee short-term resource selection

4.2.3 Pollen of popular meliponines , Africanized honeybees and lesser known species

4.2.4 Pollination ecology and population biology

4.2.5 Conclusions and ecological perspective

5. The value of plants for the Mayan stingless honey bee Melipona beecheii (Apidae: Meliponini): a pollen-based study in the Yucatán Peninsula, Mexico

5.1. Understanding the Ecology of a Mayan Resource and Cultural Icon

5.2. Baseline Studies of Invasive Honeybees and Native Neotropical Bees

5.3. Fieldwork

5.4. Pollen Analysis from Pot-Pollen Samples

5.5. Understanding Bee Resource Use in Dynamic Natural Environments

6. Melittopalynological Studies of Stingless Bees from East Coast of Peninsular Malaysia

6.1. Introduction

^len Collection by Heterotrigona itama in Tropical Island of Taman Tropika Kenyir, Terengganu

6.3. Pollen Collection and Abundance among Colonies of Lepidotrigona terminata from a Meliponary in Besut, Terengganu

6.4. Selected Flowers Producing Pollen Preferred by Stingless Bees in Terengganu

6.5. Conclusions

7. The Contribution of Palynological Surveys to Stingless Bee Conservation: a Case Study with Melipona subnitida

7.1. Introduction

7.2. Floral Resources-Dynamics: Pot-Pollen versus Pollen from the Bees' Body

7.3. Melittopalynology as Tool for Restoration Strategies: Suitable Foraging Habitats

7.4. Concluding Remarks and Future Steps

8. Pollen Storage by Melipona quadrifasciata anthidioides in a Protected Urban Atlantic Forest Area of Rio de Janeiro, Brazil

8.1. Introduction

8.2. Getting Pollen Loads and Pollen Grains by M. quadrifasciata anthidioides

8.3. Palynological Characteristics of Pollen Batches Collected from the Baskets of M. quadrifasciata anthidioides

8.3.1 Monofloral Pollen Loads

8.3.2 Bifloral Pollen Loads

8.3.3 Heterofloral Pollen Loads

8.3.4 Additional Pollen Types

8.3.5 Additional Structured Elements

8.4. Plant Families, Genera and Species Mostly Visited by M. quadrifasciata anthidioides

8.5. Conclusion

9. Angiosperm Resources for Stingless Bees (Apidae, Meliponini): A Pot-Pollen Melittopalynological Study in the Gulf of Mexico

9.1. Introduction

9.2. Background of Melittopalynological Studies in Mexico

9.3. Methods and Study Areas

9.4. Floral Resources Foraged by Melipona beecheii in the State of Campeche

9.4.1 Angiosperm Resources for Melipona beecheii

9.4.2 Physicochemical Analyses of Melipona beecheii Pot-Honey

9.5. Meliponiculture and Melitopalynological Study of Pot-Honey and Pot-Pollen in Veracruz

9.5.1 INANA’s Sustainable Meliponiculture

9.5.2 Angiosperm Resources for Scaptotrigona mexicana, Plebeia sp. and Melipona beecheii in Veracruz

9.6. Analysis of the Plant Preferences of Stingless Bees in Campeche and Veracruz, Gulf of Mexico

9.7. Angiosperms Recorded in Systematic Mexican Melittopalynological Studies of Stingless Bees

9.8. General Considerations

10. Annual Foraging Patterns of the Maya Bee Melipona beecheii (Bennett, 1831) in Quintana Roo, Mexico

10.1. Introduction

10.2. A case study

10.2.1 Field observations: registering bees activity

10.2.2. Foraging Activity to Collect Pollen and Nectar

10.2.3. Stored Pot-Honey and Pot-Pollen Reserves

10.2.4. Offspring

10.3. Correlations Between the Studied Factors<

11. Crop Pollination by Stingless Bees

11.1. Introduction

11.2. Characteristics of Stingless Bees as Pollinators

11.3. Field Crop Pollination by Stingless Bees

11.4. Greenhouse Crop Pollination by Stingless Bees

11.5. Stingless Bee Management under Greenhouse Conditions

11.6. Perspectives

12. Stingless Bees as Potential Pollinators in Agroecosystems in Argentina: Inferences from Pot-Pollen Studies in Natural Environments

12.1. Introduction

12.2. Potential Pollination by Stingless Bees: Intrinsic and Extrinsic Factors

12.2.1 Advantages and Disadvantages of a Reduced-Moderate Flight Range in Stingless Bees

12.2.2. Pollination Using Ground Nesting Stingless Bees

12.3. Pollen Spectra of Pot-Pollen in Colonies of Stingless Bees from Natural Environments

12.4. Crops Potentially Pollinated by Stingless Bees in Argentina

12.5. Spatial Variation of Crops in Argentina

12.6. Temporal Variation of Flower Availability in Agroecosystems

12.7. Crop and Non-Crop Flowerings Present in Agroecosystems Beneficial for the Maintenance of Permanent Stingless Bee Colonies

12.7.1. Pollinating the Target Crop

12.7.2. Weeds and Edge Vegetation as Complementary Flowerings for Permanent Stingless Bee Colonies

12.7.3. Diversified Agroecosystems as Best Habitat for Stingless Bee Pollination and Colony Management

12.8. Case Study: Pollination of Strawberries with Plebeia catamarcensis (Holmberg) in Santa Fe, Central Argentina

12.8.1. Strawberry Cultivation in Argentina

12.8.2 The Strawberry in Santa Fe

12.8.3 Meliponini: Potential Pollinators in Santa Fe Strawberry Crops

SECTION 2

Biodiversity, Behavior and Microorganisms of the Stingless Bees (Meliponini)

13. Stingless bees (Hymenoptera: Apoidea: Meliponini) from Gabon

13.1. Introduction

13.2. Taxonomy and Morphological Diversity of Stingless Bees in Gabon

13.3. Distribution of Stingless Bee Fauna in Gabon

13.4. Biology, Ecology and Nesting Behavior of the Stingless Bees

13.5. Knowledge and Traditional Use of Stingless Bees in Gabon

13.6. Conclusion

14. Pushing 100 Species: Meliponines (Apidae: Meliponini) in a Parcel of Western Amazonian Forest at Yasuní Biosphere Reserve, Ecuador

14.1. Yasuní Forest and Melittological Background

14.2. Discovering Meliponine Biodiversity

14.3. Species Accounts and Frequency

14.4. Insights from Comparative Morphology and other Rich Amazonian Areas

14.5. Bioprospecting for Pollination Knowledge and Sustainable Exploitation

15. Diversity of Stingless Bees in Ecuador, Pot-Pollen Standards and Meliponiculture Fostering a Living Museum for Meliponini of the World

15.1. Introduction

15.2. Megabiodiversity of Stingless Bees in Ecuador

15.3. A Revised Ecuadorian Honey Norm and Approach to Pot-Pollen Standards

15.4. Stingless Bee Keepers are Crucial for the Heritage and Conservation Mission

15.5. A Stingless Bee Window to Look at Climate Warming

15.5. Why a Living Museum to Embrace Meliponini of the World?

16. Nesting Ecology of Stingless Bees in Africa

16.1. Introduction

16.2. Meliponine Origin, Dispersal and Richness

16.3. Stingless Bee Species in Africa

16.4. Stingless Bee Nest Architecture

16.5. African Stingless Bee Nesting Behavior

17. On the Trophic Niche of Bees in Cerrado Areas of Brazil and Yeasts in their Stored Pollen

17.1. Introduction

17.2. Pollen Harvested by Native Bees of the Cerrado

17.3. Yeasts in Stored Pollen: Diversity and Ecological Role

18. A Review of the Artificial Diets Used as Pot-Pollen Substitutes

18.1. Introduction

18.1.1 Aim of the Chapter

18.1.2 How do Stingless Bees Harvest and Store their Food?

18.2. The Fermentation Process in Stingless Bees Storage Pots

18.2.1 General Characteristics of Pollen Fermentation

18.2.2 Microbial fermentation and nutritional enhancement of pollen

18.2.3 Impacts of Exogenous Compounds in Pollen

18.3. Microorganisms Present in Pot-Pollen

18.3.1 Generalities of host-associated microorganisms

18.3.2 Bacteria

18.3.3 Yeasts

18.3.4 Filamentous Fungi

18.4. Development of Artificial Diets

19. Yeast and Bacterial Composition in Pot-Pollen Recovered from Meliponini in Colombia: Prospects for a Promising Biological Resource

19.1. Introduction

19.2. General Properties of Corbicular Bee-Derived Pollen

19.3. The Key Bacterial Assemblages Known to be Associated with Bees and Pollen

19.4. Key Features of Yeast Communities Present in Pollen Collected by Bees: Recovery and Identification of Yeasts found in Pollen Collected by Four Genera of Stingless Bees from Colombia

19.5 Final remarks

SECTION 3

Stingless Bees in Culture and Traditions

20. Cultural, Psychological and Organoleptic Factors Related to the Use of Stingless Bees by Rural Residents of Northern Misiones, Argentina

20.1.      Introduction
20.2.       Southernmost Atlantic Forest Ecoregion

20.3.       Ethnobiological Field Work

20.4.       Cultural, Psychological and Organoleptic Factors Context

20.4.1     Cultural Factors

20.4.2     Psychological Factors

20.4.3   Organoleptic Factors
20.5.       Context of Exploitation of Stingless Bees

20.6.       Cultural and Psychological Factors Related to the Use of Stingless Bees

20.7.       Relationships Between Pot-Honey, Pot-Pollen, and Cultural, Psychological Factors

20.8.       Conclusions and Future Challenges

21. The Maya Universe in a Pollen Pot Native Stingless Bees in Precolumbian Maya Art
21.1.       Introduction

21.2.       Maya Bee Myths
21.3.       Small Format Modelled Sculptures

21.3.1     Small Clay Bee

21.3.2     Ceramic Beehive

21.3.3     Censers
^; Bees and Stingless Bee Keeping in a Sacred Maya Book

21.4.1   Melipona beecheii in the Tro-Cortesianus Codex

21.4.2   Hobon

21.5.       A World View of Humankind through a Bee Model as Told by a H-men, a Traditional Maya Priest
21.6.      The Maya Universe in a Pollen Pot
21.6.1     Once upon a Time, Long, Long Ago, there were Bees….

21.7.       ‘The Flower Dust’



SECTION 4

Chemical Composition, Bioactivity and Biodiversity of Pot-Pollen

22. Nutritional Composition of Pot-Pollen from Four Species of Stingless Bees (Meliponini) in South East Asia

22.1. Introduction

22.2. Shape and Volume of Stingless Bee Pollen Pots

22.3. Nutritional Composition
<22.3.1 Macronutrients of Pot-Pollen

22.3.2 Mineral Analysis

22.3.3 Fatty Acid Analysis

22.3.4 Amino Acid Analysis

22.4. Botanical Origin
22.5. Conclusions, Suggestion and Future Research

23. Characterization of Scaptotrigona mexicana Pot-Pollen from Veracruz, Mexico

23.1.       Introduction
23.2.       Traditional Values of Scaptotrigona mexicana

23.3.       Importance of Pot-Pollen in Meliponiculture <

23.4.       Palynological Analyses of Scaptotrigona mexicana Pot-pollen from Central Veracruz, Mexico

23.5.       Chemical Parameters of Scaptotrigona mexicana Pot-Pollen from Veracruz, Mexico

23.6.       Elemental Composition of Scaptotrigona mexicana Pot-pollen from Central Veracruz, Mexico
23.7.       Conclusions and Perspectives on Scaptotrigona mexicana Pot-Pollen in Mexico

24. Chemical Characterization and Bioactivity of Tetragonisca angustula Pot-Pollen from Mérida, Venezuela

24.1.      Introduction

24.2.      Proximal Analysis of Tetragonisca angustula Pot-Pollen from Mérida
24.3.       Methods to Quantify Flavonoids, Polyphenols, Proteins, and Antioxidant Activity in Ethanolic Extracts of Tetragonisca angustula Pot-Pollen

24.3.1 Preparation of Pot-Pollen Ethanolic Extracts

24.3.2 Flavonoid Content

24.3.3 Polyphenol Content

24.3.4 Protein Content
24.3.5 Antioxidant Activity by the ABTS+• Method
24.3.6   Antioxidant Activity (AOA) by Fenton-Type Reaction

24.3.7 Hydroxyl Radical Assay

24.4.      Biochemical Components and Antioxidant Activity of Ethanolic Extracts from Tetragonisca angustula Pot-Pollen 4.1   Preparation of Pot-Pollen Ethanolic Extracts

24.5.     Conclusions

25. Chemical, Microbiological and Palynological composition of the ‘Samburá’ Melipona scutellaris Pot-Pollen

25.1.       Introduction

25.2.       The ‘Samburá’ of the True “Uruçú” Bee

25.3.       Physicochemical Characteristics of ‘Samburá’

25.3.1     Moisture
25.3.2     Ash
25.3.3     Lipids

25.3.4     Protein<
25.3.5     Fiber

25.3.6     Carbohydrates

25.3.7     pH

25.3.8     Free Acidity

25.3.9     Water activity (Aw)
25.3.10 Total Energy Value
25.4.       Microbiological Characteristics of ‘Samburá’

25.5.       Pollen Analysis

26. Characterization of Pot-Pollen from Southern Venezuela
26.1.   Introduction
26.2.   Venezuelan Stingless Bees

26.3.   Botanical Origin of Venezuelan Pot-Pollen

26.4.   Proximal Analysis of Venezuelan Pot-Pollen
26.5       Bioactive Components and Antioxidant Activity of Pot-Pollen Ethanolic Extracts

26.5.1 Preparation of the Ethanolic Extracts

26.5.2 Bioactive components
26.5.3 Flavonoids by HPLC-UV
26.5.4 Antioxidant Activity of Venezuelan Pot-Pollen Homogenates

26.6.   Conclusions

27. Bioactivity and Botanical Origin of Austroplebeia and Tetragonula Australian Pot-Pollen

27.1. Introduction
27.2. Nutraceutical Properties of Bee Pollen

27.3. Botanical Origin of Australian Pot-Pollen

27.4. Flavonoids, Polyphenols and Antioxidant Activity

27.5. Antibacterial Activity of Australian Pot-Pollen

27.6. Conclusions

28. Antibacterial Activity of Ethanolic Extracts of Pot-Pollen from Eight Meliponine Species from Venezuela

28.1. Introduction

28.1.1 Biological Potential of Pollen Stored in Bee Nests

28.1.2 Aim of the Chapter

28.2. Pot-Pollen Samples and Ethanolic Extraction

28.3. Well Diffusion Agar and Minimal Inhibitory Concentration Methods

28.4. Antibacterial Activity of Venezuelan Pot-Pollen Ethanolic Extracts
28.4.1 Inhibition Zone Diameters

28.4.2 Minimal Inhibitory Concentrations

28.4.3 Antibacterial Activity of Pollen and Polyphenols

28.4. Conclusions

29. Metabolomics of Pot-Pollen from Three Species of Australian Stingless Bees (Meliponini)

29.1.       Introduction
29.1.1 Historical Accounts of Australian Pot-Pollen

29.1.2 Health Benefits of Bee Pollen

29.1.3 Botanical Sources within Flight Range<
29.1.4     Food Security

29.1.5     Research in Australian Meliponini Bee Products

29.1.6 Aim of the Chapter
29.2.       Methods of Chromatographic Analysis of Pot-Pollen

29.2.1 Sampling Pot-Pollen from Bee Hives of Australian Meliponini

29.2.2 Extraction

29.2.3 Volatiles by HS-SPME-GC-MS

29.2.4 Chemical constituents by UPLC-DAD-ESI(-)-MS/MS

29.2.5 Targeted Analyses of Pyrrolizidine Alkaloids (PA) by ESI(+)-MS/MS
29.2.6 Metabolomics of Pot-Pollen VOCs and Phenolics

29.3.      Chemometrics of Australian Meliponini Pot-Pollen

29.3.1 Volatile Organic Compounds (VOCs)

29.3.2 Secondary Metabolites by LC-UV-HRMS/MS

29.3.3 Chemometrics Using Open Source Data

29.4.      Future studies of Australian Pot-Pollen
29.4.1 Botanical and ecological studies

29.4.2 Microbiological and Metabolomics of Australian Pot-Pollen

29.4.3 Recent Trends in Australian Meliponiculture

29.5.       Conclusions

SECTION 6

Marketing and Standards of Pot-Pollen

30. Rural-Urban Meliponiculture and Ecosystems in Neotropical areas. Scaptotrigona, a Resilient Stingless Bee?
30.1. Introduction 30.2. Initiatives to Revitalize Stingless Bee Keeping

30.3. Traditional Knowledge Involving Scaptotrigona

30.4. One stingless bee, Scaptotrigona over Melipona

30.5. Naming Pot-Honey and Pot-Pollen in Labels of Commercial Products
30.6. Future of Social Interventions in Meliponiculture

31. Pot-Pollen “Samburá” Marketing in Brazil, and Suggested Legisation

31.1. Introduction
31.2. Pot-Pollen is Known as ‘Samburá’ in Brazil

31.3. Meliponíne Species Used for the Production of ‘Samburá’
31.4. Harvesting and Processing

31.5. Marketing of Meliponine ‘Samburá’
31.6. Cultural Aspects of Pot-Pollen Consumption in Brazil

31.7. Strategies to Increase the Production of ´Samburá’

31.8. Seasonality of Pot-Pollen

31.9. Pot-Pollen Production Initiatives in Brazilian States
31.10. Suggested ‘Samburá’ Standards for Pot-Pollen Legislation

Appendix A
Ethnic Names of Stingless Bees

Appendix B
Microorganisms Associated with Stingless Bees or Used to Test Antimicrobial Activity (AM)

ELECTRONIC SUPPLEMENTAL MATERIAL

Appendix C
Taxonomic Index of Bees

Appendix D
List of Bee Taxa

Appendix E
Taxonomic Index of Plant Families

Appendix F
List of Plant Taxa Used by Bees

INDEX

Professor Patricia Vit, MSc PhD

Universidad de Los Andes

Faculty of Pharmacy and Bioanalysis

Food Science Department

Mérida, Venezuela

The Sydney University

Cancer Research Group

Discipline of Biomedical Science

Sydney, Australia

Dr. Silvia R.M. Pedro

Universidade de São Paulo

School of Philosophy, Sciences and Literature

Biology Department

Brasil

Dr. David W. Roubik

Smithsonian Tropical Research Institute

Terrestrial Biology Division

Entomology Department

Panama

This book covers pot-pollen—the other product, besides honey, stored in cerumen pots by Meliponini. Critical assessment is given of stingless bee and pot-pollen biodiversity in the Americas, Africa, Asia and Oceania. Topics addressed include historical biogeography, cultural knowledge, bee foraging behavior, pollination, ecological interactions, health applications, microbiology, the natural history of bee nests, and chemical, bioactive and individual plant components in stored pollen.

Pot-pollen maintains the livelihoods of stingless bees and provides many interesting biological products that are just now beginning to be understood. The Meliponini have developed particular nesting biologies, uses of building materials, and an architecture for pollen storage. Environmental windows provide optimal temperature and availability of pollen sources for success in plant pollination and pollen storage. Palynological composition and pollen taxonomy are used to assess stingless honey bee pollination services. Pollen processing with microorganisms in the nest modifies chemical composition and bioactivity, and confers nutraceutical benefits to the honey and pollen widely relished by native people. Humans have always used stingless bees. Yet, sustainable meliponiculture (stingless bee-keeping) projects have so far lacked a treatise on pot-pollen, which experts provide in this transdisciplinary, groundbreaking volume.

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